skip to main content

Title: Drought supersedes warming in determining volatile and tissue defenses of piñon pine (Pinus edulis)

Trees are suffering mortality across the globe as a result of drought, warming, and biotic attacks. The combined effects of warming and drought onin situtree chemical defenses against herbivory have not been studied to date. To address this, we transplanted mature piñon pine trees—a well-studied species that has undergone extensive drought and herbivore-related mortality—within their native woodland habitat and also to a hotter-drier habitat and measured monoterpene emissions and concentrations across the growing season. We hypothesized that greater needle temperatures in the hotter-drier site would increase monoterpene emission rates and consequently lower needle monoterpene concentrations, and that this temperature effect would dominate the seasonal pattern of monoterpene concentrations regardless of drought. In support of our hypothesis, needle monoterpene concentrations were lower across all seasons in trees transplanted to the hotter-drier site. Contrary to our hypothesis, basal emission rates (emission rates normalized to 30 °C and a radiative flux of 1000μmol m−2s−1) did not differ between sites. This is because an increase in emissions at the hotter-drier site from a 1.5 °C average temperature increase was offset by decreased emissions from greater plant water stress. High emission rates were frequently observed during June, which were not related to plant physiological more » or environmental factors but did not occur below pre-dawn leaf water potentials of −2 MPa, the approximate zero carbon assimilation point in piñon pine. Emission rates were also not under environmental or plant physiological control when pre-dawn leaf water potential was less than −2 MPa. Our results suggest that drought may override the effects of temperature on monoterpene emissions and tissue concentrations, and that the influence of drought may occur through metabolic processes sensitive to the overall needle carbon balance.

« less
; ; ; ; ; ;
Publication Date:
Journal Name:
Environmental Research Letters
Page Range or eLocation-ID:
Article No. 065006
IOP Publishing
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    Heat and drought affect plant chemical defenses and thereby plant susceptibility to pests and pathogens. Monoterpenes are of particular importance for conifers as they play critical roles in defense against bark beetles. To date, work seeking to understand the impacts of heat and drought on monoterpenes has primarily focused on young potted seedlings, leaving it unclear how older age classes that are more vulnerable to bark beetles might respond to stress. Furthermore, we lack a clear picture of what carbon resources might be prioritized to support monoterpene synthesis under drought stress. To address this, we measured needle and woody tissue monoterpene concentrations and physiological variables simultaneously from mature piñon pines (Pinus edulis) from a unique temperature and drought manipulation field experiment. While heat had no effect on total monoterpene concentrations, trees under combined heat and drought stress exhibited ~ 85% and 35% increases in needle and woody tissue, respectively, over multiple years. Plant physiological variables like maximum photosynthesis each explained less than 10% of the variation in total monoterpenes for both tissue types while starch and glucose + fructose measured 1-month prior explained ~ 45% and 60% of the variation in woody tissue total monoterpene concentrations. Although total monoterpenes increased under combined stress, some keymore »monoterpenes with known roles in bark beetle ecology decreased. These shifts may make trees more favorable for bark beetle attack rather than well defended, which one might conclude if only considering total monoterpene concentrations. Our results point to cumulative and synergistic effects of heat and drought that may reprioritize carbon allocation of specific non-structural carbohydrates toward defense.

    « less
  2. Abstract Pinus edulis Engelm. is a short-stature, drought-tolerant tree species that is abundant in piñon-juniper woodlands throughout semiarid ecosystems of the American Southwest. P. edulis is a model species among ecophysiological disciplines, with considerable research focus given to hydraulic functioning and carbon partitioning relating to mechanisms of tree mortality. Many ecological studies require robust estimates of tree structural traits such as biomass, active sapwood area, and leaf area. We harvested twenty trees from Central New Mexico ranging in size from 1.3 to 22.7 cm root crown diameter (RCD) to derive allometric relationships from measurements of RCD, maximum height, canopy area (CA), aboveground biomass (AGB), sapwood area (AS), and leaf area (AL). Total foliar mass was measured from a subset of individuals and scaled to AL from estimates of leaf mass per area. We report a strong nonlinear relationship to AGB as a function of both RCD and height, whereas CA scaled linearly. Total AS expressed a power relationship with RCD. Both AS and CA exhibited strong linear relationships with AL (R2 = 0.99), whereas RCD increased nonlinearly with AL. We improve on current models by expanding the size range of sampled trees and supplement the existing literature for this species.more »Study Implications: Land managers need to better understand carbon and water dynamics in changing ecosystems to understand how those ecosystems can be sustainably used now and in the future. This study of two-needle pinon (Pinus edulis Engelm.) trees in New Mexico, USA, uses observations from unoccupied aerial vehicles, field measurements, and harvesting followed by laboratory analysis to develop allometric models for this widespread species. These models can be used to understand plant traits such biomass partitioning and sap flow, which in turn will help scientists and land managers better understand the ecosystem services provided by pinon pine across North America.« less
  3. Abstract
    Excessive phosphorus (P) applications to croplands can contribute to eutrophication of surface waters through surface runoff and subsurface (leaching) losses. We analyzed leaching losses of total dissolved P (TDP) from no-till corn, hybrid poplar (Populus nigra X P. maximowiczii), switchgrass (Panicum virgatum), miscanthus (Miscanthus giganteus), native grasses, and restored prairie, all planted in 2008 on former cropland in Michigan, USA. All crops except corn (13 kg P ha−1 year−1) were grown without P fertilization. Biomass was harvested at the end of each growing season except for poplar. Soil water at 1.2 m depth was sampled weekly to biweekly for TDP determination during March–November 2009–2016 using tension lysimeters. Soil test P (0–25 cm depth) was measured every autumn. Soil water TDP concentrations were usually below levels where eutrophication of surface waters is frequently observed (> 0.02 mg L−1) but often higher than in deep groundwater or nearby streams and lakes. Rates of P leaching, estimated from measured concentrations and modeled drainage, did not differ statistically among cropping systems across years; 7-year cropping system means ranged from 0.035 to 0.072 kg P ha−1 year−1 with large interannual variation. Leached P was positively related to STP, which decreased over the 7 years in all systems. These results indicate that both P-fertilized and unfertilized cropping systems mayMore>>
  4. Abstract. During the Program for Research on Oxidants: PHotochemistry, Emissions, and Transport (PROPHET) campaign from 21 July to 3 August 2016,field experiments on leaf-level trace gas exchange of nitric oxide (NO), nitrogen dioxide (NO2), and ozone (O3) were conducted for thefirst time on the native American tree species Pinus strobus (eastern white pine), Acer rubrum (redmaple), Populus grandidentata (bigtooth aspen), and Quercus rubra (red oak) in a temperate hardwood forest inMichigan, USA. We measured the leaf-level trace gas exchange rates andinvestigated the existence of an NO2 compensation point, hypothesizedbased on a comparison of a previously observed average diurnal cycle ofNOx (NO2+NO) concentrations with that simulated using amulti-layer canopy exchange model. Known amounts of trace gases wereintroduced into a tree branch enclosure and a paired blank referenceenclosure. The trace gas concentrations before and after the enclosures weremeasured, as well as the enclosed leaf area (single-sided) and gas flow rate to obtain the trace gas fluxes with respect to leaf surface. There was nodetectable NO uptake for all tree types. The foliar NO2 and O3uptake largely followed a diurnal cycle, correlating with that of the leafstomatal conductance. NO2 and O3 fluxes were driven by theirconcentration gradient from ambient to leaf internal space. The NO2 lossmore »rate at the leaf surface, equivalently the foliar NO2 deposition velocity toward the leaf surface, ranged from 0 to 3.6 mm s−1 for bigtooth aspen and from 0 to 0.76 mm s−1 for red oak, both of which are∼90 % of the expected values based on the stomatalconductance of water. The deposition velocities for red maple and white pineranged from 0.3 to 1.6 and from 0.01 to 1.1 mm s−1, respectively, and were lower than predicted from the stomatal conductance, implying amesophyll resistance to the uptake. Additionally, for white pine, theextrapolated velocity at zero stomatal conductance was 0.4±0.08 mm s−1, indicating a non-stomatal uptake pathway. The NO2compensation point was ≤60 ppt for all four tree species andindistinguishable from zero at the 95 % confidence level. This agrees withrecent reports for several European and California tree species butcontradicts some earlier experimental results where the compensation pointswere found to be on the order of 1 ppb or higher. Given that the sampledtree types represent 80 %–90 % of the total leaf area at this site, theseresults negate the previously hypothesized important role of a leaf-scaleNO2 compensation point. Consequently, to reconcile these findings,further detailed comparisons between the observed and simulated in- and above-canopy NOx concentrations and the leaf- and canopy-scaleNOx fluxes, using the multi-layer canopy exchange model withconsideration of the leaf-scale NOx deposition velocities as well asstomatal conductances reported here, are recommended.« less
  5. Abstract

    Rates of change in intrinsic water use efficiency (W) of trees relative to those in atmospheric [CO2] (ca) have been mostly assessed via short-term studies (e.g., leaf analysis, flux analysis) and/or step increases inca(e.g., FACE studies). Here we use compiled data for abundances of carbon isotopes in tree stems to show that on decadal scales, rates of change (dW/dca) vary with location and rainfall within the global tropics. For the period 1915–1995, and including corrections for mesophyll conductance and photorespiration,dW/dcafor drier tropical forests (receiving ~ 1000 mm rainfall) were at least twice that of the wettest (receiving ~ 4000 mm). The data also empirically confirm theorized roles of tropical forests in changes in atmospheric13C/12C ratios (the13C Suess Effect). Further formal analysis of geographic variation in decade-to-century scaledW/dcawill be needed to refine current models that predict increases in carbon uptake by forests without hydrological cost.